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Vocalization Behavior of the Endangered Bahama Oriole (Icterus Northropi): Ontogenetic, Sexual, Temporal, Duetting Pair, and Geographic Variation Valerie A

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3-1-2011

Vocalization Behavior of the Endangered Bahama Oriole (Icterus northropi): Ontogenetic, Sexual, Temporal, Dueting Pair, and Geographic Variation

Valerie A. Lee

Loma Linda University

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Lee, Valerie A., "Vocalization Behavior of the Endangered Bahama Oriole (Icterus northropi): Ontogenetic, Sexual, Temporal,

Dueting Pair, and Geographic Variation" (2011). Loma Linda University Electronic eses, Dissertations & Projects. 37.

htp://scholarsrepository.llu.edu/etd/37

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LOMA LINDA UNIVERSITY School of Science and Technology in conjunction with the
Faculty of Graduate Studies

____________________

Vocalization Behavior of the Endangered Bahama Oriole (Icterus northropi):
Ontogenetic, Sexual, Temporal, Duetting Pair, and Geographic Variation

by
Valerie A. Lee

____________________

A Thesis submitted in partial satisfaction of the requirements for the degree of
Master of Science in Biology

____________________

March 2011
© 2011
Valerie A. Lee
All Rights Reserved
Each person whose signature appears below certifies that this thesis in his/her opinion is adequate, in scope and quality, as a thesis for the degree Master of Science.

, Chairperson
William K. Hayes, Professor of Biology

Stephen G. Dunbar, Associate Professor of Biology Kevin E. Nick, Associate Professor of Geology

iii
ACKNOWLEDGEMENTS
I would like to thank my advisor, William K. Hayes, for getting me interested in this project back in 2005, and for all the time and support he has given me to help me through the program. Thank you to my other committee members, Stephen G. Dunbar and Kevin E. Nick; your advice and feedback have been much appreciated. Also, special thanks go to Melissa R. Price and M. Bryant J. Reynolds for their help in the field (MRP) and with the spectrographic cross-correlation analyses (MBJR). Others who helped in the field were Stephen and Vesta Myers, Keith Ingrey, Cheryl Palframan, Brian and Bev Ball, Eric Gren, Eric Robinette, and Sean Robinette. Your help was invaluable. I would like to thank my husband, James Lee, for putting up with me throughout this whole process and for helping me find easier ways to work with the data in Excel. You saved me countless hours and headaches, and I love you. Another thank you goes to my brother, Sean Robinette, for helping me with the spectrographic measurements and to learn about Raven software. Special thanks to my parents, Eric and Ann Robinette, and to my grandmothers for their love and support over the years and to God, most of all, for creating such a beautiful bird for me to study that shows me His love. Funding was provided by the Insular Species Conservation Society and by the Department of Earth and Biological Sciences at Loma Linda University. I also thank Carolyn Forbes and Aly Canestrari of Autec Navy Base for obtaining permission for us to conduct research on base, Forfar Field Station for logistical support in 2005 and 2007, and Ellen Paul of the Ornithological Council for assistance with obtaining permits. Research was approved by the Loma Linda University Institutional Animal Care and Use Committee, and conducted under a Bahamas Ministry of the Environment research permit.

iv
CONTENTS
Approval Page.................................................................................................................... iii Acknowledgements............................................................................................................ iv Table of Contents.................................................................................................................v List of Tables .................................................................................................................... vii List of Figures.................................................................................................................. viii List of Abbreviations ......................................................................................................... ix Abstract................................................................................................................................x

Chapter
1. General Introduction................................................................................................1
Ontogeny of Vocalizations ................................................................................2 Sexual Differences in Vocalizations..................................................................4 Temporal Variation............................................................................................5 Geographic Variation.........................................................................................6 Duetting..............................................................................................................6 Conservation Relevance.....................................................................................7 Objectives of Study............................................................................................8 References........................................................................................................12

2. Vocalization Behavior of the Critically Endangered Bahama Oriole
(Icterus northropi): Ontogenetic, Sexual, Temporal, and Geographic Variation ................................................................................................................18

Abstract............................................................................................................18 Introduction......................................................................................................19 Methods............................................................................................................22

Study Area .................................................................................................22 Field Work .................................................................................................24 Molecular Sex Determination....................................................................25 Vocalization Analyses ...............................................................................26 Statistical Analyses....................................................................................27

Results..............................................................................................................29

v
Molecular Sex Determination....................................................................29 Hatchling and Fledgling Vocalizations......................................................29 Adult Vocalizations ...................................................................................32 Age Variation.............................................................................................35 Differences between Sexes ........................................................................37 Time-of-Day Variation ..............................................................................39 Incubation Period Variation.......................................................................41 Geographic Variation.................................................................................41

Discussion........................................................................................................46 References........................................................................................................51

3. Duetting Behavior and Vocalizations of the Critically Endangered Bahama
Oriole (Icterus northropi) ......................................................................................56

Abstract............................................................................................................56 Introduction......................................................................................................57 Methods............................................................................................................59

Field Recording..........................................................................................59 Vocalization Analyses ...............................................................................60 Statistical Analyses....................................................................................61

Results..............................................................................................................61 Discussion........................................................................................................65 References........................................................................................................69

Appendix
Vocalization Type Spectral Characteristics .......................................................73

vi
TABLES
Tables
2.1. Spectral characters of songs from the three populations of Bahama Oriole
Page
(Icterus northropi) .................................................................................................45
3.1. Vocalization patterns, rates, and mean spectrographic cross-correlation values of duetting pairs of Bahama Orioles (Icterus northropi)............................64

3.2. Within-individual variation in spectrographic characters of vocalization bout during duetting by Bahama Oriole (Icterus northropi) pairs.........................66

vii
FIGURES

  • Figures
  • Page

1.1. An adult Bahama Oriole, Icterus northropi.............................................................9

2.1. Map of the study area, Andros Island, the Bahamas..............................................23 2.2. Representative spectrograms of hatchling (peep, chit, whine, and chu) and fledgling (chu, twitter, seet, chee) vocalizations of the Bahama Oriole (Icterus northropi) .................................................................................................31

2.3. Representative spectrograms of the five main vocalizations of the adult

Bahama Oriole (Icterus northropi), including song, whistle, chit, whine,

and squawk.............................................................................................................34
2.4. Vocalization rates of two adult age classes of the Bahama Oriole (Icterus northropi)...............................................................................................................36

2.5. Representative spectrograms from duetting male-female pairs of Bahama
Orioles (Icterus northropi).....................................................................................38

2.6. Vocalizations rates of the Bahama Oriole (Icterus northropi) at different times of the day......................................................................................................40

2.7. Vocalization rates of the Bahama Oriole (Icterus northropi) before and after the incubation period .....................................................................................42

2.8. Vocalizations rates of the Bahama Oriole (Icterus northropi) in three island populations ..................................................................................................44

3.1. Representative spectrograms from duetting male-female pairs of Bahama
Orioles (Icterus northropi).....................................................................................63

viii
ABBREVIATIONS
ANOVA ASY DFA Hz
Analyses of variance After-second-year bird or adult Discriminant function analyses Hertz, frequency
IUCN MC
International Union for Conservation of Nature Mangrove Cay

  • NA
  • North Andros

  • SA
  • South Andros

  • Sec
  • Seconds

SPCC SY
Spectrographic Cross-correlation Second-year bird or juvenile

ix

ABSTRACT OF THE THESIS

Vocalization Behavior of the Endangered Bahama Oriole (Icterus northropi):
Ontogenetic, Sexual, Temporal, Duetting Pair, and Geographic Variation

by
Valerie A. Lee
Master of Science, Graduate Program in Biology
Loma Linda University, March 2011 Dr. William K. Hayes, Chairperson

Many birds communicate via a diverse set of vocalizations, but the contexts, roles, and structure of their varied songs and calls may change with age, differ between sexes, and vary temporally and geographically. In New World orioles, most tropical species exhibit the ancestral states of sexual monochromatism (both sexes have similar plumage) and monovocalism (both sexes sing and often duet together), whereas migratory temperate species tend toward dichromatism (males brightly colored and females drab) and divocalism (males sing almost exclusively). In this study, I examined the vocalizations of the Bahama Oriole, a non-migratory, monochromatic species, to learn where it fits within this generalized dichotomy; to document sources of variation in vocalization rates and spectrographic structure; and to improve survey design for this critically endangered species. Accordingly, this study describes the primary vocalizations of the Bahama Oriole, and examines how vocalizations vary with age, between sexes, at different times of day, during the breeding season, and among the three remaining island metapopulations on Andros, The Bahamas.

x
Hatchlings and fledglings produced vocalizations that were higher pitched than those of adults. Adults possessed a large repertoire, including five main vocalization types that were delivered independently or in combination. Second-year and aftersecond-year-plumaged adults produced spectrographically similar vocalizations at similar rates. Although adult males and females could not be reliably distinguished in the field, both individuals of pairs were often heard giving the full range of vocalizations and frequently duetted together, particularly during the pre-incubation period. Antiphonal duets involved mated pairs, were limited to songs and whistle calls, and exhibited similar within-individual and between-individual variation in the spectrographic and temporal features of duets. Thus, the Bahama Oriole more closely resembles tropical oriole species (monovocal) than temperate species (divocal) in its vocalization behaviors. Adults vocalized at similar rates throughout the day prior to incubation, suggesting that surveys can be conducted at virtually any time of day during this period. Singing and most call types waned after chicks hatched, but whines increased dramatically as adults engaged in caring for their offspring. Minor but significant clinal variation in singing existed among the three metapopulations, suggesting possible cultural drift.

xi

CHAPTER ONE
GENERAL INTRODUCTION

The study of bird vocalizations can broadly inform our understanding of evolution, ecology, behavior, and conservation. Bird song serves many purposes, including species and individual recognition, mate attraction and pair bonding, territory establishment, aggression, alarm, and spacing (Langmore 1998, Marler and Slabbekoorn 2004, Rogers et al. 2006, Topp and Mennill 2007, Hall and Peters 2008). Many birds communicate via a diverse set of vocalizations, but the contexts, roles, and structure of their songs and calls may change with age, differ between the sexes, and vary temporally and geographically.
New World orioles (genus Icterus) represent an excellent group to evaluate variation in vocalizations. The molecular phylogeny of this species-rich group is well understood (Omland et al. 1999, Sturge et al. 2009). From this phylogeny, reconstruction of the evolution of plumage and song suggests that both traits are highly labile, with repeated convergence in individual elements and in overall patterns across the clade (Price et al. 2007). Both monochromatic and dichromatic species occur in this group (Hofmann et al. 2008a,b, Friedman et al. 2009), and sexual differences in singing are well documented among species, suggesting that varying levels of sexual and natural selection have contributed to the structure and roles of vocalizations in this group (Price et al. 2007, 2009).

1

Ontogeny of Vocalizations

Song development has both innate and learned properties (Slater 2003), as many species learn to vocalize at an early age from their parents and neighbors, developing dialects and personal signatures (Nowicki and Searcy 2005). Unfortunately, there is limited information on the extent to which genes, environment, or both affect song development in orioles (Rising and Flood 1998, Rising and Williams 1999, Pleasants and Albano 2001, Flood 2002, Flood et al. 2002, Brush and Pleasants 2005, Scharf and Kren 2010). Many song characters are relatively invariant within oriole taxa compared to between taxa, suggesting a strong genetic component (Price et al. 2007). However, learning may still be important to song development. Songs of hybrid Baltimore (Icterus galbula) and Bullock’s Orioles (I. bullockii) in Colorado, for example, are not intermediate, but strongly resemble those of the Baltimore Oriole, suggesting they were learned from a (Baltimore) parent rather than genetically encoded (Edinger 1985). Portions of the songs of second-year (SY) Orchard Oriole (I. spurius) males differ from those of after-second-year (ASY) males (Scharf and Kren 2010), further suggesting a learning component to oriole song development.
There is little early life history information available for most North American orioles. Information regarding the behavior and vocalizations of hatchlings, for example, is difficult to obtain due to the relative inaccessibility of oriole nests, which are tightly woven, pendulous, basket-like structures made of grass and twigs, and generally suspended from the smaller branches high in trees (Rising and Williams 1999, Pleasants and Albano 2001, Flood 2002, Flood et al. 2002, Brush and Pleasants 2005, Scharf and Kren 2010). The most detailed information on the vocalizations of young is from the

2
Baltimore Oriole (Rising and Flood 1998). Young are relatively quiet during the first week in the nest and vocalizations become louder as the hatchlings age. During the second week, they can be heard when parents approach the nest. Their calls have been described variously as teé-dee-dee, teé-dee-dee, or dee-dee-dee-dee, dee-dee-dee-dee (Bent 1958, Baumgartner and Baumgartner 1992). Just before fledging, young vocalize regularly, even between feeding visits by parents. After leaving the nest, fledglings are very noisy, uttering he-he-häe or heck-heck-he calls interspersed with low twittering (Bendire 1895). Nestlings of the Altamira Oriole (I. gularis), by comparison, were described as producing low-pitched begging calls (Brush and Pleasants 2005).
Oriole nests are frequently victimized by cowbirds, which are avian host parasites. Even less than orioles is known about the behavior and vocalizations of cowbird hatchlings and fledglings (Lowther 1993, Lowther and Post 1999, Ellison and Lowther 2009). In most oriole species, young fledge about two weeks after hatching, usually at the same time or within a two- or three-day window (Rising and Flood 1998, Rising and Williams 1999, Pleasants and Albano 2001, Flood 2002, Flood et al. 2002, Brush and Pleasants 2005, Scharf and Kren 2010). Cowbird young presumably fledge at the same time as the oriole young. Bronzed Cowbirds (Molothrus aeneus), Brown-headed

Cowbirds (Molothrus ater), and Shiny Cowbirds (Molothrus bonariensis), fledge at 10–

12 8–13, and 12–15 days, respectively (Lowther 1993, Lowther and Post 1999, Ellison and Lowther 2009).

3

Sexual Differences in Vocalizations

Among songbirds, males have traditionally been viewed as the only sex that sings.
Although female song is rare in temperate migratory songbirds and typically less complex than male song, females of many tropical non-migratory species are now known to sing complex songs (e.g., Kellner and Ritchison 1988, Gilbert and Carroll 1999, Ogden et al. 2003, Pavlova et al. 2007), often duetting with and sometimes even singing more frequently than males (Morton 1996, Price et al. 2008). This geographic difference (temporal versus tropics) is apparent in orioles (Price et al. 2008). Phylogenetic reconstruction of female song in New World blackbirds (Icteridae) suggests that malebiased song production does not result from sexual selection for complex song in males, but from selection against such songs in females (Price et al. 2009). One study suggests that female song may be the rule rather than the exception in tropical environments (Price et al. 2008). This may relate to the fact that temperate species are often dichromatic, whereas tropical species tend to be weakly dichromatic or monochromatic (Brush and Pleasants 2005). In these tropical environments, evidence suggests that females sing primarily for the same reasons as males do in temperate species (Langmore 1998, 2000, Hall 2004).
A similar reconstruction of plumage suggests that losses in female color, not elaboration of male color, has promoted the evolution of dichromatism in orioles (Hofmann et al. 2008a, Friedman et al. 2009). Thus, the traditional explanation for the evolution of sexual dimorphism, that sexual selection leads to increased male elaboration and natural selection opposes this elaboration in females (Darwin 1871, Andersson 1994), may need broad reexamination.

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  • Geographic Variation and Genetic Structure in the Bahama Oriole (Icterus Northropi), a Critically Endangered Synanthropic Species

    Geographic Variation and Genetic Structure in the Bahama Oriole (Icterus Northropi), a Critically Endangered Synanthropic Species

    Geographic variation and genetic structure in the Bahama Oriole (Icterus northropi), a critically endangered synanthropic species Melissa R. Price1,2 , Carl Person3 and William K. Hayes3 1 Department of Natural Resources and Environmental Management, University of Hawai‘i at Manoa,¯ Honolulu, HI, United States 2 Kewalo Marine Laboratory, Pacific Biosciences Research Center, University of Hawaii at Manoa, Honolulu, HI, United States 3 Department of Earth and Biological Sciences, Loma Linda University, Loma Linda, CA, United States ABSTRACT Bird species may exhibit unexpected population structuring over small distances, with gene flow restricted by geographic features such as water or mountains. The Bahama Oriole (Icterus northropi) is a critically endangered, synanthropic island endemic with a declining population of fewer than 300 individuals. It now remains only on Andros Island (The Bahamas), which is riddled with waterways that past studies assumed did not hinder gene flow. We examined 1,858 base pairs of mito- chondrial DNA sequenced from four gene regions in 14 birds (roughly 5% of the remaining population) found on the largest land masses of Andros Island (North Andros and Mangrove Cay/South Andros). We sought to discern genetic structuring between the remaining subpopulations and its relationship to current conservation concerns. Four unique haplotypes were identified, with only one shared between the two subpopulations. Nucleotide and haplotype diversity were higher for the North Andros subpopulation than for the Mangrove Cay/South Andros subpopulation. Analysis of molecular variance (AMOVA) yielded a Wright’s fixation indexF ( / of Submitted 25 August 2015 st Accepted 2 November 2015 0.60 (PFst D 0:016), with 40.2% of the molecular variation explained by within- Published 26 November 2015 population diVerences and 59.8% by among-population diVerences.
  • A Comprehensive Species-Level Molecular Phylogeny of the New World

    A Comprehensive Species-Level Molecular Phylogeny of the New World

    YMPEV 4758 No. of Pages 19, Model 5G 2 December 2013 Molecular Phylogenetics and Evolution xxx (2013) xxx–xxx 1 Contents lists available at ScienceDirect Molecular Phylogenetics and Evolution journal homepage: www.elsevier.com/locate/ympev 5 6 3 A comprehensive species-level molecular phylogeny of the New World 4 blackbirds (Icteridae) a,⇑ a a b c d 7 Q1 Alexis F.L.A. Powell , F. Keith Barker , Scott M. Lanyon , Kevin J. Burns , John Klicka , Irby J. Lovette 8 a Department of Ecology, Evolution and Behavior, and Bell Museum of Natural History, University of Minnesota, 100 Ecology Building, 1987 Upper Buford Circle, St. Paul, MN 9 55108, USA 10 b Department of Biology, San Diego State University, San Diego, CA 92182, USA 11 c Barrick Museum of Natural History, University of Nevada, Las Vegas, NV 89154, USA 12 d Fuller Evolutionary Biology Program, Cornell Lab of Ornithology, Cornell University, 159 Sapsucker Woods Road, Ithaca, NY 14950, USA 1314 15 article info abstract 3117 18 Article history: The New World blackbirds (Icteridae) are among the best known songbirds, serving as a model clade in 32 19 Received 5 June 2013 comparative studies of morphological, ecological, and behavioral trait evolution. Despite wide interest in 33 20 Revised 11 November 2013 the group, as yet no analysis of blackbird relationships has achieved comprehensive species-level sam- 34 21 Accepted 18 November 2013 pling or found robust support for most intergeneric relationships. Using mitochondrial gene sequences 35 22 Available online xxxx from all 108 currently recognized species and six additional distinct lineages, together with strategic 36 sampling of four nuclear loci and whole mitochondrial genomes, we were able to resolve most relation- 37 23 Keywords: ships with high confidence.
  • Landbird Monitoring in the Sonoran Desert Network 2012 Annual Report

    Landbird Monitoring in the Sonoran Desert Network 2012 Annual Report

    National Park Service U.S. Department of the Interior Natural Resource Stewardship and Science Landbird Monitoring in the Sonoran Desert Network 2012 Annual Report Natural Resource Technical Report NPS/SODN/NRTR—2013/744 ON THE COVER Hooded Oriole (Icterus cucullatus). Photo by Moez Ali. Landbird Monitoring in the Sonoran Desert Network 2012 Annual Report Natural Resource Technical Report NPS/SODN/NRTR—2013/744 Authors Moez Ali Rocky Mountain Bird Observatory 230 Cherry Street, Suite 150 Fort Collins, Colorado 80521 Kristen Beaupré National Park Service Sonoran Desert Network 7660 E. Broadway Blvd, Suite 303 Tucson, Arizona 85710 Patricia Valentine-Darby University of West Florida Department of Biology 11000 University Parkway Pensacola, Florida 32514 Chris White Rocky Mountain Bird Observatory 230 Cherry Street, Suite 150 Fort Collins, Colorado 80521 Project Contact Robert E. Bennetts National Park Service Southern Plains Network Capulin Volcano National Monument PO Box 40 Des Moines, New Mexico 88418 May 2013 U.S. Department of the Interior National Park Service Natural Resource Stewardship and Science Fort Collins, Colorado The National Park Service, Natural Resource Stewardship and Science office in Fort Collins, Colora- do, publishes a range of reports that address natural resource topics. These reports are of interest and applicability to a broad audience in the National Park Service and others in natural resource manage- ment, including scientists, conservation and environmental constituencies, and the public. The Natural Resource Technical Report Series is used to disseminate results of scientific studies in the physical, biological, and social sciences for both the advancement of science and the achievement of the National Park Service mission.
  • 21 Sep 2018 Lists of Victims and Hosts of the Parasitic

    21 Sep 2018 Lists of Victims and Hosts of the Parasitic

    version: 21 Sep 2018 Lists of victims and hosts of the parasitic cowbirds (Molothrus). Peter E. Lowther, Field Museum Brood parasitism is an awkward term to describe an interaction between two species in which, as in predator-prey relationships, one species gains at the expense of the other. Brood parasites "prey" upon parental care. Victimized species usually have reduced breeding success, partly because of the additional cost of caring for alien eggs and young, and partly because of the behavior of brood parasites (both adults and young) which may directly and adversely affect the survival of the victim's own eggs or young. About 1% of all bird species, among 7 families, are brood parasites. The 5 species of brood parasitic “cowbirds” are currently all treated as members of the genus Molothrus. Host selection is an active process. Not all species co-occurring with brood parasites are equally likely to be selected nor are they of equal quality as hosts. Rather, to varying degrees, brood parasites are specialized for certain categories of hosts. Brood parasites may rely on a single host species to rear their young or may distribute their eggs among many species, seemingly without regard to any characteristics of potential hosts. Lists of species are not the best means to describe interactions between a brood parasitic species and its hosts. Such lists do not necessarily reflect the taxonomy used by the brood parasites themselves nor do they accurately reflect the complex interactions within bird communities (see Ortega 1998: 183-184). Host lists do, however, offer some insight into the process of host selection and do emphasize the wide variety of features than can impact on host selection.
  • Food Habits and Feeding Behavior of the Baltimore Oriole in Costa Rica

    Food Habits and Feeding Behavior of the Baltimore Oriole in Costa Rica

    FOOD HABITS AND FEEDING BEHAVIOR OF THE BALTIMORE ORIOLE IN COSTA RICA RICHARD L. TIMKEN ALTIMORE Orioles (Zcterus galbula) are mainly insectivorous during B their summer residence in North America (Bent, 1958)) but little is known of their food habits while wintering in Central America and northern South America. Slud (1964) mentions that this bird has a varied behavior and diet while in Costa Rica, and A. F. Skutch is reported as saying that Baltimore Orioles subsist on a variety of animal and plant foods (Bent, 1958). However, no qualitative or quantitative data are available concerning the food habits and feeding behavior of this species. In Costa Rica the species occupies a wider range of habitat than do the native orioles and exceeds them in total numbers (Slud, 1964). This study was undertaken to obtain some data concerning the habits of this successful species during its stay in Costa Rica. MATERIALS AND METHODS This study was conducted at seven different sites in five of the seven Costa Rican Provinces. Collections were obtained at: Taboga, Guanacaste Province; Heredia, Heredia Province; Turrialba, Cartago Province; and San Isidro, San Jose Province. Observations of feeding behavior were obtained at the collection sites and at three other sites: San Jose, San Jose Province; San Vito, and Rincon, Puntarenas Province. Birds were collected with shotguns at different hours on several dates. Stomachs from collected specimens were removed as quickly as possible, slit and preserved in a 70 per cent ethanol solution. The number of each item was recorded per stomach and the per cent by volume of each kind of food item was estimated.